oToLITh ECo-MoRPhoLogICAL PATTERNS oF BENThIC FIShES

Thalassas, 30(1) · January 2014: 57-66
An International Journal of Marine Sciences
OTOLITH ECO-MORPHOLOGICAL PATTERNS OF BENTHIC
FISHES FROM THE COAST OF VALENCIA (SPAIN)
ANGELA M. JARAMILLO(1), ANDREA D. TOMBARI(2), VICENT BENEDITO DURA(1), MARIA EUGENI RODRIGO(3)
& ALEJANDRA V. VOLPEDO*(4,5)
(1) Grupo de Investigación Sostenibilidad Ambiental. Facultad de Ciencias Ambientales. Universidad de Ciencias Aplicadas
y Ambientales U.D.C.A. Calle 222 No.55-37. Bogotá, Colombia.
(2) Laboratorio de Biodiversidad de Vertebrados Acuáticos, Dpto. de Biodiversidad y Biologia Experimental, Facultad de Ciencias Exactas
y Naturales. Universidad de Buenos Aires, Ciudad Universitaria, Ciudad de Buenos Aires (C1428EHA) República Argentina.
(3) Instituto Agroforestal Mediterráneo. Universidad Politécnica de Valencia. Camino de Vera s/n. 46022 Valencia. España.
(4) Centro de Estudios Transdisciplinarios del Agua (CETA/INBA-CONICET), Facultad de Ciencias Veterinarias, Universidad de Buenos Aires,
Av. Chorroarín 280, Ciudad de Buenos Aires (C1427CWO) República Argentina.
(5) Instituto de Investigaciones en Producción Animal (INPA-CONICET-UBA), Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET)
*corresponding author, Alejandra V. Volpedo, Centro de Estudios Transdisciplinarios del Agua (CETA), Facultad de Ciencias Veterinarias,
Universidad de Buenos Aires, Av. Chorroarín 280, Ciudad de Buenos Aires (C1427CWO) República Argentina,
Tel/FAX: 54-11-45248484 , [email protected]
ABSTRACT
Benthic fishes exhibit different adaptive strategies and different degrees of association with the substrate. The aim of this study is to
describe the morphologic and morphometric characteristic of otolith of four species of benthonic fish off the coast of Valencia and study
the relationship between these otolith´s characteristics witch the substrate. The sagittae analysed belonged to the following species:
Scorpaena scrofa Linnaeus, 1758 (N=40), Mullus surmuletus Linnaeus, 1758 (N=29), Uranoscopus scaber Linnaeus, 1758 (N=24) and
Synaptura lusitanica Capello, 1868 (N=121). The fishes were sexed, the total length (TL) was measured in mm, and the sagittae were
removed to examine their morphology and morphometry. The E (=maximum width of the otolith (OW)/ maximum length of the otolith
(OL)%) and S (sulcus area (SS)/otolith area (OS) %) index were calculated. The E morphometric index showed a tendency towards an
elongated or circular shape and the S index showed a tendency of macula nervous to have a greater surface area of information uptake
to transmit to the fish brain. The analysis of otolith morphology and morphometry revealed the existence of different eco-morphological
patterns associated with habitat use and the type of substrate where the fish is most frequently found. The E and S index have proven to
be useful for discriminating between the sagittae of fishes from different water column uses and are associated a different substrates type.
Key words: Otoliths; Ecomorphology; Morphometry; Benthic environment.
RESUMEN
Los peces bentónicos presentan diferentes estrategias de adaptación y diferentes grados de asociación con el sustrato. El objetivo de este
estudio es describir las características morfológicas y morfométricas de otolitos de cuatro especies de peces bentónicos de la costa de
Valencia y estudiar la relación entre estas caracterisitcas de los otolitos con el sustrato. Las sagittae analizadas pertenecen a las siguientes
especies: Scorpaena scrofa Linnaeus, 1758 (N = 40), Mullus surmuletus Linnaeus, 1758 (N = 29), Uranoscopus scaber Linnaeus, 1758 (N
= 24) y Synaptura lusitanica Capello, 1868 (N = 121). Los peces fueron sexados, se midió la longitud total (lt) en mm, y las sagittae fueron
extraídas para examinar su morfología y morfometría. Se calculó el índice E (= ancho máximo de los otolitos (OW) / máxima longitud
del otolito (OL) %) y S (superficie del sulcus (SS) / superficie del otolito (OS) %). El índice morfométrico E mostró una tendencia hacia
una forma alargada o circular y el índice S mostró una tendencia de la mácula nerviosa a tener una mayor superficie de captación de
información para transmitir al cerebro del pez. El análisis de la morfología y morfometría de los otolitos reveló la existencia de diferentes
patrones eco-morfológicos asociados con el uso del hábitat y el tipo de sustrato donde los peces se encuentran con mayor frecuencia. Los
índices E y S han demostrado ser útiles para discriminar entre las sagittae de peces que hacen un uso diferencial de la columna de agua
y que están asociados a diferentes tipos de sustratos.
Palabras clave: otolitos, ecomorfología, morfometría, ambiente bentónico.
Thalassas, 30(1) · January 2014
57
ANGELA M. JARAMILLO, ANDREA D. TOMBARI, VICENT BENEDITO DURA, MARIA EUGENI RODRIGO & ALEJANDRA V. VOLPEDO
Figure 1.
Figure 1: Study area.
Introduction
Benthic fishes exhibit different adaptive strategies
and different degree of association with the substrate.
This association mainly depends on the type of substrate
and on the morphology, physiology and behaviour of the
fish (Lindsey 1978; Menni 1983).
In general, fishes living on soft substrate (slimy
and sandy substrates) burrow partially in the bottom
while stalking prey. Most of these predators, such as
the Pleuronectiformes, are characterised by rapid and
short-distance movements. Fishes found on hard substrate
(rocky substrate) hide in holes or stand motionless,
waiting for the prey to approach. These are represented
by the Scorpaeniformes and Serranids, and other fish
families.
The differences in habitat use and the different
communication strategies of fishes may also affect
different structures as the otolith sagittae (Cruz and
Lombarte 2004; Lombarte and Cruz 2007; Lombarte et
al. 2010). Otoliths are three paired of calcified structures
(sagittae, lapilli and asterici) found in the inner ear, used
for balance and/or hearing in all teleost fishes. The otolith
shape is species-specific. Environmental factors (Aguirre
and Lombarte 1999; Torres et al. 2000; Gauldie and
Crampton 2002; Volpedo and Echeverría 2003; Volpedo
et al. 200; Volpedo and Fuchs 2010), ontogeny (Tombari
et al. 2005; Gonzalez Naya et al. 2012), physiology as
58
Thalassas, 30(1) · January 2014
the hearing capabilities associated with specialization in
acoustic communication (Popper and Fay 1993; Paxton
2000; Lombarte and Cruz 2007) and the phylogeny
(Nolf and Tyler 2006), could affect the morphology,
the morphometry and the microstructure of sagittae
(Volpedo and Fernández Cirelli 2006; Volpedo et al.
2007). The literature studying the morphometric variation
of the otoliths of benthic fish are limited (Volpedo and
Echeverría 2003).
The aim of this study is to describe the morphologic
and morphometric characteristic of otolith of four
23 of Valencia
species of benthonic fish off the coast
(Scorpaena scrofa Linnaeus, 1758, Mullus surmuletus
Linnaeus, 1758, Uranoscopus scaber Linnaeus, 1758, and
Synaptura lusitanica Brito Capello, 1868), and study the
relationship between these otolith´s characteristics witch
the substrate.
Materials and Methods
The analysis was based on the sagittae of four fish
species: Scorpaena scrofa (N=40), Mullus surmuletus
(N=29), Uranoscopus scaber (N=24) and Synaptura
lusitanica (N=121). Fish samples were mature specimens
and present the typical morphological pattern of their
otoliths. Between July 2004 and March 2005, fishes were
caught in Bay of Cullera (39º 12’- 38º 59’ N and 0º 09’- 0º
15’W) (Fig. 1) with gillnets operated by commercial, and
artisanal fishermen.
Figure 2
OTOLITH ECO-MORPHOLOGICAL PATTERNS OF BENTHIC FISHES FROM THE COAST OF VALENCIA (SPAIN)
Figure 2:
Otolith morphology of sagittae. Antirostrum (AR), cauda (C), crenulated rim (CR), excisura (Ex), horseshoe-shaped area depression (HAD), irregular rim (IR),
ostium (Os), regular rim (RR), sulcus (S) and ventral area depresion (VAD).
Samples were kept at -18ºC until processing in
the laboratory. The specimens were identified using
appropriate keys (Corbera et al. 2000; Whitehead et al.
2001; Froese and Pauly 2008).
Fishes were sexed, their total length (TL) was measured to the nearest mm, and the sagittae were removed
from the otic capsules. They were prepared for examination and drawing, and measured under stereoscopic
microscope to the nearest 0.1 mm.
Otolith morphological description is based on the
terminology proposed by Volpedo and Echeverría (2000)
and Tused et al. (2008). The following morphological
characters were considered in medial face: geometrical
shape, type of rim, presence of rostrum, antirostrum
and excisura, type of sulcus, presence of ostium and
cauda and presence de area depression. In addition, the
topography of the lateral face (side opposite the sulcus)
was described (Fig. 2).
The maximum width of the otolith (OW) and
maximum length of the otolith (OL) were measured in the
medial face of each right sagitta (Fig. 3).
OL
OS
The E and S index were calculated. The E index (E =
OW/OL%) was calculated to assess if the otoliths showed
a tendency towards an elongated or circular shape. The S
index is (SS)/(OS) %). The otolith and sulcus areas were
measured using a digital image processing system. The
OL*100/fish size was calculated, total length (TL, mm)
was the reference for fish size. The E index and OL*100/
TL were calculated from 45 species from otolith images
provided AFORO data base (http://aforo.cmima.csic.es)
purposes comparatives.
The statistical analysis was based on the mean values
of the different parameters. The Student’s t-test (Sokal
and Rolf 1987; Zar 2004) was used to compare between
the length and width of the right and left sagittae and
between the sagittae of males and females.
A ANCOVA was used to test for significant
differences in the E and S index among species, followed
by the Tukey’s multiple comparison test. Linear regression
analysis was performed to determine the relationship
between each morphometric character (OW and OL) and
fish total length (TL), and the regression parameters and
the coefficient of determination (R 2) were calculated.
Statistical analysis was performed using SPSS Inc., 16.0
for Windows and StatGraphics Centurion XV, version
15.2.06.
Results
Morphological features of sagittae
OW
SS
Figure 3:
Morphometry of otoliths. Maximum length of the otolith (OL), maximum
width of the otolith (OW), otolith surface (OS), sulcus surface (SS).
The sagitta of Scorpaena scrofa is elongated, with
a regular dorsal rim and an irregular ventral rim. The
rostrum is conspicuous and represents 28% approximately
of otolith length. The excisura and antirostrum are
conspicuous. The sulcus is divided into a funnel-shaped
24 rim
ostium and a cauda slightly directed to the ventral
distally. A ventral area depression is parallel to the sulcus.
The medial face is convex and the lateral face is concave
with radiating grooves (Fig. 4 A).
Thalassas, 30(1) · January 2014
59
ANGELA M. JARAMILLO, ANDREA D. TOMBARI, VICENT BENEDITO DURA, MARIA EUGENI RODRIGO & ALEJANDRA V. VOLPEDO
Figure 4.
A
B
C
D
E
Figure 4:
Morphology of sagittae. A: Scorpaena scrofa. B: Mullus surmuletus, C: Uranoscopus scaber, D: left sagitta of Synaptura lusitanica,
E: right sagitta of Synaptura lusitanica. Scale: 2 mm
In Mullus surmuletus the sagitta is oval, with
scalloped or crenulated ventral and dorsal rims; the
rostrum represents 30% approximately of otolith length
and the excisura has a notch. The ostium is perforated. The
sulcus is open anteriorly and posteriorly, is surrounded
by ventral and dorsal ridges, and divided into ostium
and cauda. The ostium is funnel-shaped and the cauda
is initially straight and then curves distally towards the
ventral rim. It has dorsal and ventral area depressions,
oriented parallel to the sulcus. The medial face is convex
and the lateral face is concave and shows radiating
grooves. (Fig. 4 B)
The sagitta of Uranoscopus scaber is oval, with
regular rims. It lacks rostrum, antirostrum or excisura.
The sulcus is closed anteriorly and posteriorly and not
divided into ostium and cauda. The area depression is
absent. The medial and lateral faces are slightly flat, with
the latter showing radiating grooves (Fig. 4 C).
The left and right sagittae of Synaptura lusitanica
60
Thalassas, 30(1) · January 2014
show morphological differences. The left sagitta is
oblong, with regular rims. There is a quadrangularshaped process at the postero-dorsal tip.
26 The rostrum is
poorly developed and the excisura and antirostrum are
short. The sulcus is open anteriorly, ostium and cauda
are differentiated. The cauda is small and almost circular
in the posterior part of the sulcus, and is surrounded
by a horseshoe-shaped area depression. The medial
face is convex and the lateral face is undulated (Fig. 4
D). The right sagitta is oblong, with irregular rims and
a projection at the postero-dorsal tip. The ostium and
cauda are differentiated. The collum is present between
the ostium and the cauda. The horseshoe-shaped area
depression is present. The medial face is convex and the
lateral face is undulated (Fig. 4 E).
Morphometrical features of otoliths
Table 1 shows the significance of the differences in
length and width between the right and left sagittae, as
determined by the Student’s t-test.
58
48
OTOLITH ECO-MORPHOLOGICAL PATTERNS OF BENTHIC FISHES FROM THE COAST OF VALENCIA (SPAIN)
38
S.scrofa
Figure 5
M.surmuletus
U.scaber S.lusitanica
A
Species
98
Hard sustrate Mixed sustrate
Soft sustrate
60
Sp1
50
E
78
45
68
40
Sp3
35
Sp2
25
20
S.scrofa
M.surmuletus
U.scaber S.lusitanica
15
A
10
Species
60
Sp1
S 30
48
38
Soft sustrate
Mixed sustrate
Sp3
55 Sp2
88
58
Hard sustrate
Hard sustrate
55 Sp2
Mixed sustrate
5
Soft sustrate
0
Sp3
S.scrofa
M.surmuletus
U.scaber S.lusitanica
B
Species
50
Figure 5:
Mean45
values and standard deviation. A) E index (E= OW/OL%). B) S index (S= SS/ OS %). sp1: species associated to soft substrate (Table 4), sp2: species
40
associated to hard substrate (Table 4), sp3: species associated to mixed substrate (Table 4).
35
In relation to fish relative Sp1
size (OL*100/TL), the
S 30
otoliths
of S. scrofa and U. scaber were the largest
25 and 3.03-4.83, respectively), while those of M.
(4.11-4.64
surmuletus
and S. lusitanica were the smallest (2.70-3.48
20
and 2.26-3.11,
respectively).
15
10
The right and left sagittae of M. surmuletus, S. scrofa
and U.5 scaber no showed morphological differences in
B
0
S.scrofa
M.surmuletus
S.lusitanica
the topography
of the inner
and outerU.scaber
face of
the otolith.
S. lusitanica present differences
Species between the right and left
sagitta (Fig. 4).
The results of the comparison between the sagittae
of males and females are shown in Table 2. The otoliths
of females of M. surmuletus, S. lusitanica and U. scaber
were significantly larger than those of males of a similar
length, while the otoliths of S. scrofa did not differ
significantly between sexes.
The regression parameters (a and b) and coefficient of
determination (R2) for the relationships between the morphometric characters of the sagittae and fish total length are
presented in Table 3. The regressions were significant for
all species (Table 3) and data from symmetric fishes fitted
well to the linear regression model (R2>0.80). The lowest
values of R2 were obtained for S. lusitanica, particularly
for the relationship between otolith width and fish total
length. In this species showing morphologically different
right and left sagittae, the values of R2 calculated from the
right sagittae were higher than those from the left sagittae.
The lowest and highest values of the E morphometric
index were obtained for S. scrofa (40.3%) and M. sur-
muletus (70.54%), respectively, while S. lusitanica and
U. scaber showed similar values (55.63 % and 53.12 %,
respectively) (Fig. 5 A). ANCOVA showed significant differences in the E morphometric index among the studied
species (F(3;216) = 117.2; P < 0.05), and the Tukey’s multiple
comparison test revealed significant differences between
M. surmuletus, S. scrofa and the other species. There were
no significant differences in the E morphometric index
between S. lusitanica and U. scaber.
The mean value of S indexes for M. surmuletus
are highest (26.12 % ± 4.57) and lowest for S. scrofa
(11.58% ± 3.73). S. lusitanica and U. scaber showed
intermediate values (12.94% ± 3.55 % and 18.74% ±
8.32%, respectively) (Figure 5 B).
27
ANCOVA showed significant differences in the S
index among studies species (F(3;216) = 107.3; P < 0.05) and
the Tukey’s multiple comparison test revealed significant
differences between M. surmuletus, S. scrofa and the other
species. There were no significant differences in the S
morphometric index between S. lusitanica and S. scrofa.
Discussion
The otoliths of benthic fishes from the coast of
Valencia showed different eco-morphological patterns,
which could be associated with the type of substrate
where the fish is most frequently found and habitat use.
In order to relate our results with those of the
Otolith AFORO Database and have a reference in the
interpretation of these, we have obtained the E index and
Thalassas, 30(1) · January 2014
61
Table 1. Morphometric features (mean ± SD) of sagitta in the studied species and t-student
ANGELA M. JARAMILLO, ANDREA D. TOMBARI, VICENT BENEDITO DURA, MARIA EUGENI RODRIGO & ALEJANDRA V. VOLPEDO
results for comparison between right and left otolith. N. sample number, TL= fish total
Table
1:
legth; OL = maximum length of the otolith;
OW=
maximum width of the otolith. *P>0.05
Morphometric features (mean ± SD) of sagitta in the studied species and t-student results for comparison between right and left otolith. N. sample number,
significative
TL= fish total legth; OL = maximum length of the otolith; OW= maximum width of the otolith. *P>0.05 significative
Species
S. scrofa
M. surmuletus
U. scaber
S. lusitanica
TL
Morphometric
Right Otolith
Left Otolith
(mm)
features
(mm)
(mm)
224 ± 7
OL
9.8 ± 1.6
OW
205 ± 4
234 ± 5
235 ± 4
t-value
N
P
9.1 ± 2.1
0.87
24
0.4
3.8 ± 0.6
3.6 ± 0.8
0.34
OL
6.3 ± 0.8
6.3 ± 0.8
0.054
OW
4.5 ± 0.5
4.4 ± 0.5
0.705
OL
9.2 ± 2.1
9.2 ± 2.1
0.087
OW
4.9 ± 0.9
4.8 ± 0.9
0.097
OL
6.3 ± 1.0
5.9 ± 1.1
2.56
OW
3.4 ± 0.5
3.7 ± 0.8
2.91
0.7
20
0.9
0.5
24
0.9
0.9
121
< 0.05*
< 0.05*
substrates is 79.09% ±8.37 and S 20.13 ± 12.7 %. In the
case of species associated with hard substrates have an
average of E index of 48.47% ±5.23 and S 35.43±20.11 %.,
while the 10 species associated both to mixed substrates
have an average of 56.20% ±5.81 and S 34.63±23.6 %.
the relative sizes of the otoliths in relation to the TL, from
45 species of western Mediterranean, north and central
eastern Atlantic fishes (Tuset et al. 2008; Otolith AFORO
Database 2012). The data are shown in table 4.
In table 4, of the 45 species, 18 species inhabit soft
species
of of
Bothidae,
substrates,
17 hard
substrates andbetween
10 speciesmorphometric
are associated features The
Table
2. Comparison
(mean
± SD)
sagitta Cynoglossidae,
of both sex Gobiidae
and Soleidae families generally present otolith with the E
to mixed substrates (hard and soft) (Fischer et al. 2007).
index
valuesOW=
>60%maximum
and are associated
soft substrates
The average
of the E index
of species
withlength
soft of the
and t-student
results.
OL =associated
maximum
otolith;
width ofto the
otolith. *P>0.05 significative
Table 2:
Comparison between morphometric features (mean ± SD) of sagitta of both sex and t-student results. OL = maximum length of the otolith;
OW= maximum width of the otolith. *P>0.05 significative.
Species
Morphometric
Female
Male
features
(mm)
(mm)
OL
9.8 ± 1.7
OW
S. scrofa
M. surmuletus
U. scaber
S. lusitanica
62
Thalassas, 30(1) · January 2014
t-values
g.l.
P
9.7 ± 1.6
0.17
24
0.87
3.8 ± 0.6
4.0 ± 0.8
0.98
OL
6.7 ± 0.6
5.8 ± 0.5
3.1
OW
4.7 ± 0.5
4.2 ± 0.5
2.23
OL
9.9 ± 2.2
9.1 ± 1.2
3.01
OW
5.2 ± 1.0
4.4 ± 0.6
2.9
OL
6.3 ± 1.1
5.7 ± 0.7
3.4
OW
3.7 ± 0.7
3.3 ± 0.5
4.03
16
0.33
20
< 0.05*
< 0.05*
40
< 0.05*
< 0.05*
121
< 0.05*
< 0.05*
Table 3. Regression linealOTOLITH
parameters
pf morphometrical features of sagitta in relation to
ECO-MORPHOLOGICAL PATTERNS OF BENTHIC FISHES FROM THE COAST OF VALENCIA (SPAIN)
total length of fish. a: intercept, b: slope, R2: determination coefficient. LOL. left otolith
length, LOW left otolith width, OL: right length
otolith, OW: right otolith width, TL. Total
Table 3:
Regression lineal parameters pf morphometrical features of sagitta in relation to total length of fish. a: intercept, b: slope, R2: determination coefficient. LOL.
length of
fish.
left otolith length, LOW left otolith width, OL: right length otolith, OW: right
Specie
otolith width, TL. Total length of fish.
a
b
R2
TL vs OL
3.40
0.26
0.87
TL vs OW
1.17
0.11
0.88
TL vs OL
2.76
0.17
0.81
TL vs OW
1.17
0.13
0.84
TL vs OL
-0.43
0.41
0.87
TL vs OW
0.30
0.19
0.88
TL vs OL
0.36
0.25
0.85
TL vs LOL
-0.02
0.25
0.62
TL vs OW
1.15
0.09
0.40
TL vs LOW
0.83
0.12
0.25
Morphometrical
features
S. scrofa
M. surmuletus
U. scaber
S. lusitanica
and S index values between 10.38-53.16 %. The species of
Muraenidae, Scorpaenidae, Serranidae, Trachinidae and
Tripterygiidae families present otolith with the E index
values <60% and S index values between 13.25-32.91 and
are associated to hard substrates. The labrids, mullids and
sparids present species associated to soft substrates and
hard substrates (Table 4). Xyrichthys novacula (labrids)
present an E index value of 89.41% inhabiting soft substrate, while the labrids associated to hard substrate (Table
4) present E index values between 50.00-55.11%. Mullus
barbatus barbatus present an E index value of 75.96%
inhabinting sof substrates (Fischer et al. 2007). The sparid
species of soft bottom (Dentex macrophthalmus) present E
index values of 80.46 %, and sparid species of hard bottom
(Dentex dentex) E index values of 57.44% (Table 4).
The fishes burrowing into soft bottom substrates (U.
scaber and S. lusitanica), have oblong otoliths of variable
size in relation to fish total length and an E morphometric
index with 53.12% and 55.63%. This species inhabit sandy
or muddy bottoms between 10 to 90 m deep, feeding off
small fish, crustaceans, molluscs and polychaetes (Calvin
2000, Jaramillo 2009). U. scaber and S lusitanica shared
similar habitat, however the latter is a trophic specialist,
predating predominantly on polychaetes (Jaramillo 2009).
As the E and the S indexes are similar it can be suggested
that otolith morphology and morphometry reflected
the similarities between U. scaber and S lusitanica
habitat characteristics and behavior. These relationships
established for other species of different habitats (Volpedo
and Echeverria 2003; Volpedo et al. 2008; Cruz and
Lombarte 2004; Lombarte and Cruz 2007).
The otolith of S. lusitanica shows a horseshoe-shaped
area depression, which results from cranial remodelling
and eye migration during larval metamorphosis. This
feature is shared by other flatfishes such as Achiropsetta
tricholpis Norman, 1930, Etropus longimanus Norman,
1933, Paralichthys orbynianus (Valenciennes, 1839),
P.isosceles Jordan, 1891, P. brasiliencies (Ranzani, 1842),
P. patagonicus Jordan, 1889 and Xystreuris rasile Jordan,
1891 (Volpedo and Echeverría 1997).
S. scrofa is present on hard bottoms, and M. surmuletus
is found on mixed bottoms (soft and hard substrates), both
species showed a different morphological pattern in their
otoliths and water column used. S. scrofa18
is a solitary
fish common on hard substrate and in caves, and it may
move to sandy bottom sediments despite its sedentary
behaviour (Corbera et al. 2000). The otolith of this species
is 4.5% of fish total length, with an E index of 40.3%, and
the S index (11.58%) is the lowest value of studied species
in this paper. The otolith of S. scrofa is elongated shape
and well-developed rostrum are also present in other
species of the genus such as S. elongata, S. maderensis,
S. notata and S. porcus (Tuset et al. 2008), and in hardbottom serranid and notothenid fishes (Volpedo and
Echeverría 2003).
Thalassas, 30(1) · January 2014
63
ANGELA M. JARAMILLO, ANDREA D. TOMBARI, VICENT BENEDITO DURA, MARIA EUGENI RODRIGO & ALEJANDRA V. VOLPEDO
ANGELA M. JARAMILLO, ANDREA D. TOMBARI, VICENT BENEDITO DURA, MARIA EUGENI RODRIGO & ALEJANDRA V. VOLPEDO
Table 4. E index and OL/TL relation in species of literature.1-soft substrate, 2: hard
4:
substrate,3: mixed bottoms (soft and hardTable
substrates).
Table 4:
E index and OL/TL relation in species of literature.1-soft substrate, 2: hard substrate,3: mixed bottoms (soft and hard substrates).
E index and OL/TL relation in species of literature.1-soft substrate, 2: hard substrate,3: mixed bottoms (soft and hard substrates).
Family
E%
(OL*100)/TL SS/OS% Substrate
Arnoglossus imperialis (Rafinesque, 1810)
69.70
1.886
30.493
1
Arnoglossus laterna (Walbaum, 1792)
66.67
2.371
24.691
1
Arnoglossus rueppelii (Cocco, 1844)
67.77
2.305
24.585
1
Arnoglossus thori Kyle, 1913
64.94
3.138
53.165
1
Bothus podas (Delaroche, 1809)
66.93
1.434
28.097
1
Callionymus maculatus Rafinesque, 1810
47.80
2.661
54.762
3
Callionymus risso Lesueur, 1814
49.86
2.823
53.126
3
Synchiropus phaeton (Günther, 1861)
47.81
2.314
20.408
3
Cepolidae
Cepola macrophthalma (Linnaeus, 1758)
55.10
2.305
20.719
3
Citharidae
Citharus linguatula (Linnaeus, 1758)
61.60
3.035
17.580
3
Cynoglossidae
Symphurus ligulatus (Cocco, 1844)
80.72
3.557
12.032
1
Symphurus nigrescens Rafinesque, 1810
87.17
1.965
15.741
1
Deltentosteus quadrimaculatus (Valenciennes, 1837)
84.22
4.444
10.578
1
Gobius niger Linnaeus, 1758
78.04
4.549
10.384
1
Lesueurigobius friesii (Malm, 1874)
87.27
4.650
10.601
1
Mullidae
Mullus barbatus barbatus Linnaeus, 1758
75.96
2.176
18.636
1
Soleidae
Bathysolea profundicola (Vaillant, 1888)
85.51
2.070
14.38
1
Monochirus hispidus Rafinesque, 1814
85.51
2.070
24.27
1
Pegusa lascaris (Risso, 1810)
91.47
2.413
13.62
1
Solea senegalensis Kaup, 1858
82.66
2.464
19.72
1
Solea solea (Linnaeus, 1758)
79.19
1.700
13.61
1
Lithognathus mormyrus (Linnaeus, 1758)
59.91
3.531
20.303
3
Dentex macrophthalmus (Bloch, 1791)
80.46
4.894
14.090
1
Trachinus draco Linnaeus, 1758
47.58
4.935
17.045
2
Trachinus radiatus Cuvier, 1829
48.62
3.828
19.153
2
Coris julis (Linnaeus, 1758)
55.11
1.957
18.135
2
Labrus merula Linnaeus, 1758
50.00
1.726
14.250
2
Labrus viridis Linnaeus, 1758
51.35
1.930
14.620
2
Symphodus cinereus (Bonnaterre, 1788)
62.79
1.911
7.210
3
Symphodus doderleini Jordan, 1890
55.02
2.544
16.080
3
Symphodus mediterraneus (Linnaeus, 1758)
56.60
2.136
9.434
3
Symphodus ocellatus (Linnaeus, 1758)
65.58
1.925
7.339
3
Symphodus rostratus (Bloch, 1791)
60.09
2.027
6.874
2
Symphodus tinca (Linnaeus, 1758)
56.13
1.843
5.893
2
Xyrichthys novacula (Linnaeus, 1758)
89.41
2.040
18.750
1
Muraenidae
Muraena helena Linnaeus, 1758
44.25
0.718
32.915
2
Scorpaenidae
Helicolenus dactylopterus (Delaroche, 1809)
50.64
4.542
23.552
2
Scorpaena elongata Cadenat, 1943
40.30
5.448
16.071
2
Scorpaena notata Rafinesque, 1810
43.12
5.508
21.278
2
Scorpaena porcus Linnaeus, 1758
42.96
4.656
13.252
2
Epinephelus marginatus (Lowe, 1834)
48.57
2.714
28.820
2
Serranus cabrilla (Linnaeus, 1758)
42.31
4.160
23.909
2
Serranus scriba (Linnaeus, 1758)
43.54
3.388
30.073
2
57.44
3.186
13.985
2
44.37
1.603
17.620
2
Bothidae
Callionymidae
Gobiidae
Sparidae
Trachinidae
Labridae.
Serranidae
64
64
Specie
Thalassas, 30(1) · January 2014
Sparidae
Thalassas, 30(1)
· January 2014 Dentex dentex (Linnaeus, 1758)
Tripterygiidae
Tripterygion delaisi Cadenat & Blache, 1970
19
OTOLITH ECO-MORPHOLOGICAL PATTERNS OF BENTHIC FISHES FROM THE COAST OF VALENCIA (SPAIN)
M. surmuletus mainly inhabits mixed bottoms (rocky
bottoms and soft substrates), and undergoes vertical
movements between 5 and 100 m in depth (Froese and
Pauly 2008). This species showed the highest value of E
and S indexes (70.54% and 26.13%, respectively), which
may be explained by a differential use of the water column.
The indexes values obtained for M. surmuletus are similar
at de values determinate by Volpedo et al. (2008) in
mesopelagic Antarctic fishes that make extensive vertical
migrations and other Mediterranean fishes species that
using soft and hard substrates (table 4).
The high value of S index could be associated with
water column uses, this could be a support for adaptive
physiological features as specialization in acoustic
communication in deep waters in order to compensate
the reduction of light with depth (Lombarte and Cruz
2007), among other factors. The otolith morphology of
this species (geometric shape, type of rims, presence of
rostrum, and type of sulcus) is similar to that of other
species of the same genus such as M. barbatus, M.
mullus and M. argentinae Hubbs, Marini & Hubbs, 1933
(Volpedo and Echeverría 2000; Tuset et al. 2008). The
differences in otolith morphometry observed between
males and females of M. surmuletus, S. lusitanica and U.
scaber were also found for Prionotus nudigula Ginsburg,
1950 (Volpedo and Thompson 1998). This result may be
due to the fact that these species are sexually dimorphic
in size, with females growing at a lower rate and being
larger than males (Reñones et al. 1995; Jaramillo 2009).
According to Volpedo and Echeverría (2003), the
differences in the E and S index among fish species
may be due not only to phylogenetic factors but also to
environmental and bioecological factors. In this work, the
E and S index have proven to be useful for discriminating
between the sagittae of fishes from different substrate. On
this basis, it can be seen as a valuable tool for studies of
trophic ecology in the coast of Valencia.
This work contributed to the knowledge of the
bioecology of commercially important benthic fishes and
provided key information for studying the trophic ecology
of fish-eating species and fishery management.
Acknowledgments
We are grateful to the “Conselleria de Territori I
Habitatge. Generalitat Valenciana”, to the Universidad
de Buenos Aires (UBACYT 20620110100007), to the
National Scientific and Technical Research Council
(CONICET) and ANPCYT (PICT N° 2010-1372) for the
financial support to this project. The authors grateful to
C. Assis for critical revision of manuscript and for their
comments, and G.E. Comte for the technical support in
images analyzed.
References
Aguirre H, Lombarte A (1999). Ecomorphologic comparisons of
sagittae in Mullus barbatus and M. surmuletus. Journal of
Fish Biology, 55: 105-114.
Calvin JC (2000) El ecosistema marino mediterráneo guía de
su flora y fauna 2°Ed. Juan Carlos Calvin Calvo, Murcia,
España, 797 pp.
Corbera J, Sabatés A, García-Rubies A (2000). Peces de Mar
de la Península Ibérica. Guía de Campo. Ed. Planeta,
Barcelona. 75 pp.
Cruz A, Lombarte A (2004). Otolith size and its relationship
with colour pattern and sound production. Journal of Fish
Biology, 65: 1512–1525.
Fischer ST, Patzner RA, Müller CHG, Winkler HM (2007).
Studies on the ichthyofauna of the coastal waters of Ibiza
(Balearic Islands, Spain). Rostocker Meeresbiologische
Beiträge 18: S. 30-62
Froese R, Pauly D. (eds.) (2008). FishBase. World Wide Web
electronic publication. www.fishbase.org, version (02/2008).
Gauldie RW, Crampton JS (2002). An ecomorphological
explication of individual variability in the shape of the fish
otolith: comparison of the otolith of Hoplostethus atlanticus
with other species by depth. Journal of Fish Biology, 60:
1221–1240.
Gonzalez Naya MJ, Tombari A, Volpedo AV, Gómez SE (2012).
Size related changes in sagitta otoliths of Australoheros
facetus (Pisces; Cichlidae) from South America. Journal of
Applied Ichthyology, 725-755
Jaramillo A (2009). Estudio de la biología trófica de cinco
especies de peces bentónicos de la costa de Cullera.
Relaciones con la acumulación de metales pesados. Ph.D.
thesis, Univ. Politécnica Valencia.
Lindsey CC (1978). Form, function, and locomotory habits in
fish. In: Hoar WS, Randall DJ. (Eds.) Fish physiology. Vol.
II. Academic Press, New York, 100 pp.
Lombarte A, Cruz A (2007). Otolith size trends in marine fish
communities from different depth strata. Journal of Fish
Biology, 71 (1): 53-76.
Lombarte A, Palmer M, Matallanas J, Gómez-Zurita J, MoralesNin B (2010). Ecomorphological trends and phylogenetic
inertia of otolith sagittae in Nototheniidae. Environmental
Biology of Fishes, 89:607-618.
Menni RC (1983). Los peces en el medio marino. Estudio Sigma
SRL. Argentina, 171 pp.
Nolf D, Tyler H (2006). Otolith evidence concerning
interrelationship of caproid zeiform and tetradontiform
fishes. Bulletin de l’Institut Royal des Sciences Naturelles
de Belgique, 76: 147-189.
Paxton JR (2000). Fish otoliths: do sizes correlate with taxonomic
group, habitat and/ or luminescence? Philosophical
Thalassas, 30(1) · January 2014
65
Transactions of the Royal Society B: Biological Sciences
355: 1299–1303.
Popper AN, Fay RR (1993). Sound detection and processing by
fish: critical review and major research questions. Brain
Behavior and Evolution, 41: 14-38.
Reñones O, Massutí E, Morales-Nin B (1995). Life history of red
mullet (Mullus surmuletus) from the bottom-trawl fishery
off the island of Mallorca (NW Mediterranean). Marine
Biology, 123: 411-419.
Sokal RR, Rolf FL (1987). Introduction to biostatistics, 2nd
Edition. Freeman, New York, 887 pp.
Tombari AD, Volpedo AV, Echeverría DD ( 2005). Desarrollo
de la sagitta en juveniles y adultos de Odontesthes
argentinensis (Valenciennes, 1835) y Odontesthes
bonariensis (Valenciennes, 1835) de la provincia de Buenos
Aires, Argentina (Teleostei: Atheriniformes). Revista
Chilena de Historia Natural, 78:623-633.
Torres GJ, Lombarte A, Morales-Nin (2000). Variability of
the sulcus acusticus in the sagittal otolith of the genus
Merluccius (Merluccidae). Fisheries Research, 46: 5–13.
Tuset VM, Lombarte A, Assis CA (2008). Otolith atlas for the
Western Mediterranean, North and Central Eastern Atlantic.
Scientia Marina, 72 (S1): 7-198.
Volpedo AV, Echeverría DD (1997). Morfología de la sagitta de
lenguados del Mar Argentino (Pisces: Pleuronectiformes).
Thalassas, 13: 113-126.
Volpedo AV, Thompson GA (1998). Diferencias en el crecimiento
de las sagittae de Prionotus nudigula Ginsburg, 1950
(Pisces: Triglidae) en relación al sexo. Boletín del Instituto
Español de Oceanografía, 12(1): 3-16.
Volpedo AV, Echeverría DD (2000). Catálogo y claves de otolitos
para la identificación de peces del mar argentino. 1. Peces
de importancia económica. Dunken, Buenos Aires, 88 pp.
Volpedo AV, Echeverría DD ( 2003). Ecomorphological patterns
of the sagitta in fish on the continental shelf off Argentine.
Fisheries Research, 60: 551-560.
Volpedo AV, Fernández Cirelli A ( 2006). Otolith chemical
composition as a useful tool for sciaenids stock discrimination
in Southwestern Atlantic. Scientia Marina, 70 (2): 325-334.
Volpedo AV, Miretzky P, Fernández Cirelli A (2007). Stocks
pesqueros de Cynoscion guatucupa y Micropogonias
furnieri de la costa atlántica de Sudamérica: comparación
entre métodos de identificación. Memoria de la Fundacion
La Salle de Ciencias Naturales, 165: 115-130.
Volpedo AV, Tombari AD, Echeverría DD (2008). Ecomorphological patterns of the sagitta of Antarctic fish. Polar
Biology, 31 (5): 635-640.
Volpedo AV, Fuchs DV (2010). Ecomorphological patterns of
the lapilli of Paranoplatense Siluriforms (South America).
Fisheries Research, 102: 160-165.
Whitehead PJP, Bauchot ML, Hureau JC, Nielson J, Tortonese
T (Eds) (2001). Fishes of the north-eastern Atlantic and
the Mediterranean. World Biodiversity Database CD-ROM
Series. ETI UNESCO, Amsterdam.
Zar J (2004). Biostatistical Analysis. Fourth Edition. Prentice
Hall International, INC. New Jersey, USA, 663 pp.
(Received: July, 15, 2013; Acepted: October, 24, 2013)
66
Thalassas, 30(1) · January 2014